Showing papers in "Zoologica Scripta in 2011"

A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera)

Abstract: Zahiri, R., Kitching, I. J., Lafontaine, J. D., Mutanen, M., Kaila, L., Holloway, J. D. & Wahlberg, N. (2010). A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera). —Zoologica Scripta, 40, 158–173. To examine the higher level phylogeny and evolutionary affinities of the megadiverse superfamily Noctuoidea, an extensive molecular systematic study was undertaken with special emphasis on Noctuidae, the most controversial group in Noctuoidea and arguably the entire Lepidoptera. DNA sequence data for one mitochondrial gene (cytochrome oxidase subunit I) and seven nuclear genes (Elongation Factor-1α, wingless, Ribosomal protein S5, Isocitrate dehydrogenase, Cytosolic malate dehydrogenase, Glyceraldehyde-3-phosphate dehydrogenase and Carbamoylphosphate synthase domain protein) were analysed for 152 taxa of principally type genera/species for family group taxa. Data matrices (6407 bp total) were analysed by parsimony with equal weighting and model-based evolutionary methods (maximum likelihood), which revealed a new high-level phylogenetic hypothesis comprising six major, well-supported lineages that we here interpret as families: Oenosandridae, Notodontidae, Erebidae, Nolidae, Euteliidae and Noctuidae.

The phylogeny and limits of Elateridae (Insecta, Coleoptera): is there a common tendency of click beetles to soft-bodiedness and neoteny?

Abstract: Kundrata, R. & Bocak, L. (2011). The phylogeny and limits of Elateridae (Insecta, Coleoptera): is there a common tendency of click beetles to soft-bodiedness and neoteny? —Zoologica Scripta, 40, 364–378. Phylogenetic relationships in Elateroidea were investigated using partial 18S and 28S rDNA and rrnl and cox1 mtDNA sequences with special interest in the phylogeny of Elateridae and the position of soft-bodied lineages Drilidae and Omalisidae that had been classified as families in the cantharoid lineage of Elateroidea until recently. Females in these groups are neotenic and almost completely larviform (Drilidae) or brachypterous (Omalisidae). The newly sequenced individuals of Elateridae, Drilidae, Omalisidae and Eucnemidae were merged with previously published datasets and analysed matrices include either 155 taxa with the complete representation of fragments or 210 taxa when some fragments were missing. The main feature of inferred phylogenetic trees was the monophyly of Phengodidae + Rhagophthalmidae + Omalisidae + Elateridae + Drilidae with Omalisidae regularly occupying a basal node in the group; Drilidae were embedded as a terminal lineage in the elaterid subfamily Agrypninae and soft-bodied Cebrioninae were a part of Elaterinae. The soft-bodied males and incompletely metamorphosed females originated at least three times within the wider Elateridae clade. Their atypical morphology has been considered as a result of long evolutionary history and they were given an inappropriately high rank in the previous classifications. The frequent origins of these modifications seem to be connected with modifications of the hormonal regulation of the metamorphosis. The superficial similarity with other soft-bodied lineages, such as Cantharidae, Lycidae, Lampyridae, Phengodidae and Rhagophthalmidae is supposed to be a result of homoplasious modifications of the ancestral elateroid morphology due to the incomplete metamorphosis. The results of phylogenetic analyses are translated in the formal taxonomic classification. Most Drilidae are placed in Elateridae as a tribe Drilini in Agrypninae, whilst Pseudeuanoma and Euanoma are transferred from Drilidae to Omalisidae. The subfamily Cebrioninae is placed in Elaterinae as tribes Cebrionini and Aplastini. Oxynopterini, Pityobiini and Semiotini are lowered from the subfamily rank to tribes and classified in Denticollinae.

Integrative inventory of Bolivia’s lowland anurans reveals hidden diversity

Abstract: Jansen, M., Bloch, R., Schulze, A. & Pfenninger, M. (2011). Integrative inventory of Bolivia’s lowland anurans reveals hidden diversity. —Zoologica Scripta, 40, 567–583. We conducted an integrative species inventory and re-evaluated the taxonomic status of anuran populations from lowland Bolivia that we putatively ordered to 59 a priori species. The analyses of own and published data of mtDNA (16S), bioacoustics, and adult and tadpole morphology, however, revealed a total number of 69 divergent lineages. With the data at hand, the taxonomic assignment of only 27 from 69 lineages (39.1%) to nominal species was supported, the assignment of nine lineages (14.5% of 69) was attributed to incertae sedis, and another 33 divergent lineages (47.8%) could not yet be assigned to any species. Of those lineages, 16 were confirmed as divergent to other species by at least two of the following lines of evidence: high genetic divergence, strong differences in advertisement calls, morphology or ecology. These ‘confirmed genealogical lineages’ are suggested to belong to good species following standards of divergence of the group under study, many of them probably being candidates for formal descriptions. Another 17 lineages were considered as ‘unconfirmed genealogical lineages’ showing significant difference to all other described species only in one line of evidence. Within 6 of our 69 revealed lineages, we detected deep conspecific lineages. Some species in this study showed remarkable intraspecific call variation not congruent with genetic divergence. Our results suggest that many Bolivian anuran species that are assigned to supposedly ‘widespread’ South American species apparently belong to previously unrecognized lineages with supposedly smaller distribution areas. Our data on a regional scale urge for an integrative taxonomic review of these and other widespread lowland Neotropical anuran species, as many of them may actually represent species complexes. However, a complete inventory of Neotropical amphibians apparently is a far more complex challenge than previously thought, and only cooperative, supraregional and cross-country approaches will reveal the actual amphibian diversity in the Neotropics. We postulate that Bolivia’s lowlands can be assessed as a region joining distinct regional faunal diversity and thus deserve reinforced conservation and research efforts to fulfil its unique situation as climatic and biogeographic transition zone.

Phylogeny of a trans‐Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single early colonization of Australia

Abstract: Heinicke, M. P., Greenbaum, E., Jackman, T. R. & Bauer, A. M. Phylogeny of a trans-Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single early colonization of Australia. —Zoologica Scripta, 40, 584–602. The genus Gehyra (34 species) is rare among squamate reptile radiations in spanning continents, extending from southeast Asia to Australia and Polynesia. Among the family Gekkonidae sensu stricto, Gehyra is the only genus that is species rich in Australia. We performed molecular phylogenetic, divergence timing, and ancestral area analyses to investigate the evolutionary and biogeographic history of Gehyra. Phylogenetic analyses resolve Hemiphyllodactylus as the closest relative of Gehyra. Some data also link Perochirus to this group, but previously suggested relationships with other morphologically similar genera of geckos are not supported. Within Gehyra, three geographically discrete clades are recovered, respectively, concentrated in Asia, the Pacific islands and Australia. Ancestral area analyses suggest that Gehyra originated in Asia, with a single colonization of Australia occurring in the mid-Cenozoic. This date places the time of Gehyra colonization prior to those of other Australian gekkonid geckos, but after the near-endemic pygopodoid geckos, a Gondwanan relictual group. Based on these dates, times of origin may best explain relative differences in species diversity among Australian gekkotans. In contrast, although originating earlier, Gehyra is less diverse in Asia than in Australia. This pattern may be explained by the long-term presence of many competing, ecologically similar genera in Asia (e.g. Gekko, Hemidactylus, Lepidodactylus), whereas nearly all pygopodoids in Australia (the only gekkotans present at the time of colonization of Australia by Gehyra) are ecologically distinct.

Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia)

Abstract: Kuntner, M., May-Collado, L. J. & Agnarsson, I. (2010). Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia). —Zoologica Scripta, 40, 1–15. Phylogenies play an increasingly important role in conservation biology providing a species-specific measure of biodiversity – evolutionary distinctiveness (ED) or phylogenetic diversity (PD) – that can help prioritize conservation effort. Currently, there are many available methods to integrate phylogeny and extinction risk, with an ongoing debate on which may be best. However, the main constraint on employing any of these methods to establish conservation priorities is the lack of detailed species-level phylogenies. Afrotheria is a recently recognized clade grouping anatomically and biologically diverse placental mammals: elephants and mammoths, dugong and manatees, hyraxes, tenrecs, golden moles, elephant shrews and aardvark. To date, phylogenetic studies have focused on understanding higher level relationships among the major groups within Afrotheria. Here, we provide a species-level phylogeny of Afrotheria based on nine molecular loci, placing nearly 70% of the extant afrotherian species (50) and five extinct species. We then use this phylogeny to assess conservation priorities focusing on the widely used evolutionary distinctiveness and global endangeredness (EDGE) method and how that compares to the more recently developed PD framework. Our results support the monophyly of Afrotheria and its sister relationship to Xenarthra. Within Afrotheria, the basal division into Afroinsectiphilia (aardvark, tenrecs, golden moles and elephant shrews) and Paenungulata (hyraxes, dugongs, manatees and elephants) is supported, as is the monophyly of all afrotherian families: Elephantidae, Procaviidae, Macroscelididae, Chrysochloridae, Tenrecidae, Trichechidae and Dugongidae. Within Afroinsectiphilia, we recover the most commonly proposed topology (Tubulidentata sister to Afroscoricida plus Macroscelidea). Within Paenungulata, Sirenia is sister to Hyracoidea plus Proboscidea, a controversial relationship supported by morphology. Within Proboscidea, the mastodon is sister to the remaining elephants and the woolly mammoth sister to the Asian elephant, while both living elephant genera, Loxodonta and Elephas are paraphyletic. Top ranking evolutionarily unique species always included the aardvark, followed by several species of elephant shrews and tenrecs. For conservation priorities top ranking species always included the semi-aquatic Nimba otter shrew, some poorly known species, such as the Northern shrew tenrec, web-footed tenrec, giant otter shrew and Giant golden mole, as well as high profile conservation icons like Asian elephant, dugong and the three species of manatee. Conservation priority analyses were broadly congruent between the EDGE and PD methodologies. However, for certain species EDGE overestimates conservation urgency as it, unlike PD, fails to account for the status of closely related, but less threatened, species. Therefore, PD offers a better guide to conservation decisions.

Phylogeny of haplo–diploid, fungus-growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data

Abstract: Cognato, A. I., Hulcr, J., Dole, S. A. & Jordal, B. H. (2010). Phylogeny of haplo-diploid, fungus-growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data. —Zoologica Scripta, 40, 174–186. The ambrosia beetle tribe Xyleborini currently contains 30 genera and approximately 1200 species which are distributed throughout worldwide forests with most diversity located in the tropics. They also represent the most invasive scolytines in North America. Despite economic concerns and biological curiosity with this group, a comprehensive understanding of generic boundaries and the evolutionary relationship among species is lacking. In this study, we include 155 xyleborine species representing 23 genera in parsimony and Bayesian analyses using 3925 nucleotides from mitochondrial (COI) and nuclear genomes (28S, ArgK, CAD, EF-1α) and 39 morphological characters. The phylogenies resulting from the parsimony analyses, which treated gap positions either as missing or fifth character states, and the Bayesian analysis were generally similar. Clades with high support or posterior probabilities were found in all trees, while those with low support were not recovered by all analyses. Fourteen of the 23 genera were monophyletic although not all relationships among the genera were resolved. We show monophyly of several species groups associated with particular morphological and biological characters and suggest recognition of these groups as genera. Most interesting was the monophyly of South and Central American species representing several genera. This finding suggests recent and fast radiation of xyleborines in the New World accompanied by morphological and biological diversification.

The use of the DNA barcode gene 16S mRNA for the clarification of taxonomic problems within the family Sertulariidae (Cnidaria, Hydrozoa)

Abstract: Moura, C. J., Cunha, M. R., Porteiro, F. M. & Rogers, A. D. (2011). The use of the DNA barcode gene 16S mRNA for the clarification of taxonomic problems within the family Sertulariidae (Cnidaria, Hydrozoa). —Zoologica Scripta, 40, 520–537. The Sertulariidae are the most speciose family of the Hydrozoa, and their systematics are problematic. Here, 77 new 16S mtDNA sequence ‘barcodes’ of Atlantic and Mediterranean sertulariids were subject to phylogenetic analyses along with another 29 already deposited in Genbank. Amongst the 12 nominal genera and around 40 putative species analysed, several taxonomic inconsistencies and corroborations were highlighted by the genetic data. Patterns of morphological characters between the specimens analysed genetically were extremely congruent with molecular estimates. The genera Sertularia, Dynamena and Symplectoscyphus were found to be polyphyletic, and the genus Sertularella paraphyletic. The validity of Sertularella robusta at a species level was confirmed, along with the exclusion of Sertularelloides cylindritheca from the genus Sertularella. Potential cryptic species were detected within Sertularella fusiformis, Sertularella mediterranea, Sertularella ellisii, Sertularia distans, Dynamena disticha, Thuiaria articulata and Diphasia margareta. Conversely, probable cases of synonomy were revealed for the species pairs: Sertularia cupressina and Sertularia argentea, Amphisbetia minima and Amphisbetia operculata, and Thuiaria thuja and T. articulata. 16S haplotype data also revealed probable phylogeographic structure across localities of the NE Atlantic and Mediterranean. As an alternative ‘DNA barcode’, the 16S gene has the great advantage of being easily amplified and sequenced across hydrozoan taxa. Furthermore, the marked increase in deposited 16S sequences in public databases facilitates the identification of hydrozoan samples to the level of family, genus, and especially species or even populations, in a manner that is coherent with phenotypic and geographic/environmental differences.

Rethinking the evolution of extant sub-Saharan African suids (Suidae, Artiodactyla)

Abstract: Although African suids have been of scientific interest for over two centuries, their origin, evolution, phylogeography and phylogenetic relationships remain contentious. There has been a long-running debate concerning the evolution of pigs and hogs (Suidae), particularly regarding the phylogenetic relationships among extant Eurasian and African species of the subfamily Suinae. To investigate these issues, we analysed the mitochondrial and nuclear DNA sequences of extant genera of Suidae from Eurasia and Africa. Molecular phylogenetic analyses revealed that all extant sub-Saharan African genera form a monophyletic clade separate from Eurasian suid genera, contradicting previous attempts to resolve the Suidae phylogeny. Two major sub-Saharan African clades were identified, with Hylochoerus and Phacochoerus grouping together as a sister clade to Potamochoerus. In addition, we find that the ancestors of extant African suids may have evolved separately from the ancestors of modern day Sus and Porcula in Eurasia before they colonised Africa. Our results provide a revision of the intergeneric relationships within the family Suidae.

An integrative taxonomic revision of the Cape Verdean skinks (Squamata, Scincidae)

Abstract: Miralles, A., Vasconcelos, R., Perera, A., Harris, D. J. & Carranza, S. (2010). An integrative taxonomic revision of the Cape Verdean skinks (Squamata, Scincidae). —Zoologica Scripta, 40, 16–44. A comprehensive taxonomic revision of the Cape Verdean skinks is proposed based on an integrative approach combining (i) a phylogenetic study pooling all the previously published molecular data, (ii) new population genetic analyses using mitochondrial and nuclear data resulting from additional sampling, together with (iii) a morphological study based on an extensive examination of the scalation and colour patterns of 516 live and museum specimens, including most of the types. All Cape Verdean species of skinks presently recognised, formerly regarded as members of the genera Mabuya Fitzinger, 1826 and Macroscincus Bocage, 1873 are considered as members of the Cape Verdean endemic genus Chioninia Gray, 1845. The new phylogeny and networks obtained are congruent with the previously published phylogenetic studies, although suggesting older colonization events (between 11.6 and 0.8 Myr old), and indicate the need for taxonomic changes. Intraspecific diversity has been analysed and points to a very recent expansion of Chioninia delalandii on the southern islands and its introduction on Maio, to a close connection between Chioninia stangeri island populations due to Pleistocene sea-level falls and to a generally low haplotypic diversity due to the ecological and geological characteristics of the archipelago. Three new consistent morphological synapomorphies supporting two of the four main clades of the genus have been identified. The complex taxonomic status of Euprepes fogoensis O’Shaughnessy, 1874 has been resolved and a lectotype has been designated for this species; Chioninia fogoensis nicolauensis (Schleich, 1987) is elevated to species rank, whereas Chioninia fogoensis antaoensis (Schleich, 1987) is now regarded as a junior subjective synonym of C. fogoensis. Additionally, one new subspecies of Chioninia vaillanti and two of Chioninia spinalis are described (Chioninia vaillanti xanthotis ssp. n., Chioninia spinalis santiagoensis ssp. n. and Chioninia spinalis boavistensis ssp. n.) and a lectotype has been designated for Mabuia spinalis Boulenger, 1906. Finally, an identification key for the Chioninia species is presented.

Refuting the six-genus classification of Penaeus s.l. (Dendrobranchiata, Penaeidae): a combined analysis of mitochondrial and nuclear genes

Abstract: The taxonomic revision in 1997 of the shrimps formerly classified in Penaeus s.l. has been one of the most controversial issues on systematics of the decapods in recent years. Since Perez Farfante & Kensley (Penaeoid and Sergestoid Shrimps and Prawns of the World, 1997) split this long-accepted taxon into six genera, much debate has been devoted to their proposed new classification scheme; this has taken place because there are serious doubts whether the said scheme could reflect the evolutionary relationships among the 29 Penaeus s.l. species. Although these shrimps can be easily separated into several groups morphologically, whether these subdivisions are truly monophyletic and warrant a generic rank continues to be hotly debated among taxonomists. This study examined a total of 2425 bp sequences from three nuclear protein genes (enolase, phosphoenolpyruvate carboxykinase and sodium–potassium ATPase α-subunit), and the mitochondrial 16S and 12S rRNA gene of 18 Penaeus s.l. shrimps and 13 other species in the family Penaeidae. Our phylogenetic analyses strongly support the monophyly of Penaeus s.l. and, concurring with previous studies that used the mitochondrial genes alone, the paraphyly of both Penaeus s.s. (sensu Perez Farfante & Kensley, Penaeoid and Sergestoid Shrimps and Prawns of the World, 1997) and Melicertus, rendering them non-natural groupings. Our study reveals two lineages: Penaeus s.s. + Fenneropenaeus + Litopenaeus + Farfantepenaeus and Melicertus + Marsupenaeus, which exhibit genetic divergences comparable with those among other penaeid genera. However, all the morphological characters, which are emphasized by Perez Farfante and Kensley and used to separate Penaeus s.l., do not correlate with the grouping revealed by the present, perhaps decisive, phylogenetic result. Such disparity may arise from selection on the morphology of genitalia and convergent evolution. Our molecular data clearly refute the six-genus classification, and we advocate the restoration of the old Penaeus genus (=Penaeus s.l.) definition which is the only classification scheme with both the morphological and the molecular data being in agreement.

Cenozoic mystery birds – on the phylogenetic affinities of bony‐toothed birds (Pelagornithidae)

Abstract: Mayr, G. (2011) Cenozoic mystery birds – on the phylogenetic affinities of bony-toothed birds (Pelagornithidae). —Zoologica Scripta, 40, 448–467. The extinct Cenozoic bony-toothed birds (Pelagornithidae) are characterized by the occurrence of unique spiky projections of the osseous jaws and are among the most distinctive neornithine taxa. Earlier authors considered these marine birds to be most closely related to ‘Pelecaniformes’ or Procellariiformes, but recent phylogenetic analyses resulted in a sister group relationship to Anseriformes. This latter hypothesis was, however, coupled with a non-monophyly of galloanserine or even neognathous birds, which is not supported by all other current analyses. The character evidence for anseriform affinities of pelagornithids is thus reassessed, and it is detailed that the alleged apomorphies cannot be upheld. Pelagornithids lack some key apomorphies of galloanserine birds, and analysis of 107 anatomical characters did not support anseriform affinities, but resulted in a sister group relationship between Pelagornithidae and Galloanseres. By retaining a monophyletic Galloanseres, this result is in better accordance with widely acknowledged hypotheses on the higher-level phylogeny of birds. The (Pelagornithidae + Galloanseres) clade received, however, only weak bootstrap support, and some characters, such as the presence of an open frontoparietal suture, may even support a position of Pelagornithidae outside crown-group Neognathae.

Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa)

Abstract: Fritz, U., Branch, W. R., Hofmeyr, M. D., Maran, J., Prokop, H., Schleicher, A., Siroký, P., Stuckas, H., Vargas-Ramirez, M., Vences, M. & Hundsdorfer, A. K. (2010). Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa). —Zoologica Scripta, 40, 115–125. With 18 currently recognised species, Pelusios is one of the most speciose chelonian genera worldwide, even though the taxonomy of some species is contentious. Recent investigations suggested that the closely related, but morphologically distinct genus Pelomedusa is paraphyletic with respect to Pelusios, and that Pelomedusa consists of nine deeply divergent lineages. Using three mitochondrial and three nuclear DNA fragments (2054 bp mtDNA, 2025 bp nDNA), we examined for the first time the phylogeny of Pelusios by molecular means. Our analyses included all Pelusios species, except the probably extinct P. seychellensis, as well as the nine Pelomedusa lineages. The results showed that Pelusios and Pelomedusa are reciprocally monophyletic. Limited sampling of Pelusios species and homoplasy introduced by remote outgroups most likely explain the paraphyly of Pelomedusa in previous studies. The distinctiveness of most Pelusios species was confirmed, but none of the currently recognised species groups within Pelusios was monophyletic. In Pelusios rhodesianus and P. sinuatus distinct genetic lineages were discovered, suggestive of cryptic taxa. In contrast, the recognition of the weakly differentiated P. castaneus and P. chapini as full species is doubtful, as is the validity of the Malagasy and Seychellois subspecies of P. castanoides. GenBank sequences of P. williamsi were nested within P. castaneus, but the morphological distinctiveness of the two species makes it likely that the GenBank sequences (derived from a turtle from the pet trade) are misidentified. Divergence among the distinct genetic lineages of Pelomedusa equals or exceeds the differences among Pelusios species, supporting the view that Pelomedusa is a species complex.

Molecular prospecting for cryptic species in Phyllodistomum lacustri (Platyhelminthes, Gorgoderidae)

Abstract: Rosas-Valdez, R., Choudhury, A. & Perez-Ponce de Leon, G. (2011). Molecular prospecting for cryptic species in Phyllodistomum lacustri (Platyhelminthes, Gorgoderidae). —Zoologica Scripta, 40, 296–305. Partial sequences of the 28S ribosomal RNA and cytochrome c oxidase I (COI) genes were compared among populations of a widely distributed and morphologically uniform digenean species in North America, Phyllodistomum lacustri, a parasite characteristically associated with ictalurid catfishes. Specimens were collected from the urinary bladder of ictalurid hosts in six localities of North America, spanning most of the latitudinal range of this freshwater fish group. Sequences of other congeneric species, including a potentially close relative, P. staffordi, were also obtained and used for comparison. Analyses of both molecular markers show very low or no intrapopulation variation within each sampling site. However, samples of P. lacustri from different hosts and regions exhibit varying levels of interpopulation genetic differences. Such differences are explained by the wide geographical distribution and host range of the ictalurids they parasitize, particularly in ictalurids distributed in Mexican freshwaters where they experienced a diversification process. Variation in both nuclear and mitochondrial genes and phylogenetic analyses, in conjunction with geographical and host information (drainage isolation and endemicity of the host species), indicate that at least three populations show potential as cryptic species.

Molecular phylogeny of venus clams (Mollusca, Bivalvia, Veneridae) with emphasis on the systematic position of taxa along the coast of mainland China

Abstract: Chen, J., Li, Q., Kong, L. & Zheng, X. (2011). Molecular phylogeny of venus clams (Mollusca, Bivalvia, Veneridae) with emphasis on the systematic position of taxa along the coast of mainland China. —Zoologica Scripta, 40, 260–271. Veneridae is the most richly speciose family of heterodont bivalves with high ecological and economic value. Attention to the Veneridae systematics has been raised since traditional conchology-based ideas on relationships among the venerids were challenged by recent studies using molecular makers and other new approaches and methods. Herein, DNA sequence information from fragments of two mitochondrial genes (COI and 16S) and one nuclear protein-coding gene (H3) for 135 taxa (128 venerids, five nonvenerid veneroids and two other outgroups) are used to reconstruct the phylogenetic relationships of venus clams under maximum parsimony, maximum likelihood and Bayesian inference approaches. According to our molecular results, the traditional Veneridae is not recovered as monophyletic and most of the nominal subfamilies and genera formed para-polyphyletic clades. The findings indicate that the current venerid classification cannot validly reflect a natural subdivision. In the present study, the classification of taxa along the coast of mainland China within this family are also revised based on their phylogenetic position and morphological characters. The synonymization of chionine genus Placamen with Clausinella is rejected. Chionine subgenera Anomalodiscus and Cryptonema are given full generic rank again and incorporated into Venerinae and Tapetinae, respectively. Tapetine Marcia hiantina, M. japonica and M. marmorata were distantly related to Katelysia spp., so assigning those three species into the genus Katelysia by some malacologists is rejected herein. Our results also evidence that the synonymization of the genus Tigammona and Periglypta might be inappropriate.

Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae)

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Molecular barcoding of skates (Chondrichthyes: Rajidae) from the southern Northeast Atlantic

Abstract: Serra-Pereira, B., Moura, T., Griffiths, A. M., Gordo, L. S. & Figueiredo, I. (2010). Molecular barcoding of skates (Chondrichthyes: Rajidae) from the southern Northeast Atlantic. —Zoologica Scripta, 40, 76–84. Due to their vulnerability to fishing pressure, many species of skate (Rajidae) in the Northeast Atlantic are undergoing declines in abundance. The assessment of stock status and subsequent proposal of management measures are often complicated by high levels of species diversity and endemism, coupled with morphological and ecological conservatism, which makes distinguishing between species difficult. To improve the identification of skates and investigate the phylogenetic position of endemic species the cytochrome c oxidase subunit I (COI) was sequenced in 12 species (Dipturus oxyrinchus, Leucoraja naevus, Leucoraja circularis, Neoraja iberica, Raja brachyura, Raja clavata, Raja maderensis, Raja microocellata, Raja miraletus, Raja montagui, Raja undulata, Rostroraja alba) inhabiting the Portuguese waters. Based on sequence divergence R. maderensis and R. clavata only differ by 1% of the 652 bp COI sequence, questioning the recognition of R. maderensis (considered to be endemic to Madeira and the Azores), as a reproductively isolated species. Otherwise, there was clear phylogenetic support for the different genera and all the remaining species, although the genetic divergence was low compared to other chordates. In particular, COI analysis allowed clear identification of the morphologically similar species R. brachyura and R. montagui.

Phylogeny of the highly divergent zoanthid family Microzoanthidae (Anthozoa, Hexacorallia) from the Pacific

Abstract: Fujii, T. & Reimer, J. D. (2011). Phylogeny of the highly divergent zoanthid family Microzoanthidae (Anthozoa, Hexacorallia) from the Pacific. —Zoologica Scripta, 40, 418–431. In this study, one new family, one new genus and two new species of zoanthids from rubble zones spanning the temperate, subtropical and tropical Pacific Ocean are described. Two new species are described, Microzoanthus occultus sp. n. and Microzoanthus kagerou sp. n., both belonging to the new genus Microzoanthus and new family Microzoanthidae, and they can be clearly distinguished both morphologically and genetically from each other and other zoanthids by their very small size, reduced or absent stolon, habitat usually on the bottom side of rubble zone rocks, and divergent and distinct DNA (cytochrome oxidase subunit I, mitochondrial 16S ribosomal DNA, internal transcribed spacer region of ribosomal DNA) sequences. The phylogenetic analyses clearly show Microzoanthidae fam. n. to be genetically far different from all other hexacorallians at the order level, but the macrocnemic arrangement of mesenteries and other morphological characters (colonial specimens with narrow stolons, two rows of tentacles sand encrustation) clearly place these specimens within the order Zoantharia. This study demonstrates how it is highly likely the existence of many marine invertebrate taxa remains overlooked, and that widely distributed groups such as Microzoanthidae fam. n. remain to be discovered.

Phylogeny of macrophagous leeches (Hirudinea, Clitellata) based on molecular data and evaluation of the barcoding locus

Abstract: IDDALLSubmitted: 5 August 2010Accepted: 23 November 2010doi:10.1111/j.1463-6409.2010.00465.xOceguera-Figueroa, A., Phillips, A. J., Pacheco-Chaves, B., Reeves, W. K. & Siddall, M. E.(2010). Phylogeny of macrophagous leeches (Hirudinea, Clitellata) based on molecular dataand evaluation of the barcoding locus. — Zoologica Scripta, 00, 000–000.The phylogenetic relationships of macrophagous leech species are studied using two mito-chondrial [cytochrome c oxidase subunit I (COI) and 12S rDNA] and two nuclear (28SrDNA and 18S rDNA) markers. The complete dataset analysed in this study included 49terminals and 5540 aligned characters. Phylogenetic analyses were performed under twooptimality criteria: Maximum Parsimony and Maximum Likelihood. The monophyly ofthe two currently recognized families (i.e. Erpobdellidae and Salifidae) is confirmed andwell supported. The phylogenetic position of Gastrostomobdellidea is studied for the firsttime and found to be sister to family Salifidae nested well within Erpobdelliformes. Previ-ously recognized taxonomic arrangements were evaluated and discarded through successiveconstraint analyses. Correlation between morphology and phylogeny was notable in Salifi-dae but not in Erpobdellidae. Variability of COI, the barcoding locus, was examined acrossspecies leading to the recognition of the invasive Barbronia weberi in Mexico, Costa Rica,Germany, South Africa and Taiwan.Corresponding author: Alejandro Oceguera-Figueroa, American Museum of Natural History,Division of Invertebrate Zoology, Central Park West at 79th Street, New York, NY 10024, USA.E-mail: aoceguera@amnh.orgAlejandro Oceguera-Figueroa and Anna J. Phillips, Department of Biology, Graduate Center, TheCity University of New York and Division of Invertebrate Zoology, American Museum of NaturalHistory, Central Park West at 79th street, New York, NY 10024, USA. E-mail: ajphillips@amnh.orgBernald Pacheco-Chaves, Escuela de Biologi´a, Universidad de Costa Rica, Co´digo Postal: 11501-2060, San Jose´, Costa Rica. E-mail: bernald.pacheco@gmail.comWill K. Reeves, US Army Public Health Command Region – Pacific Unit 45006, MCHB-AJ-TLD APO, AP 96343-5006, USA. Public Health Command, US Army Japan. E-mail: will.reeves@us.army.milMark E. Siddall, Sackler Institute for Comparative Genomics, American Museum of Natural His-tory, Central Park West at 79th Street, New York, NY 10024, USA. E-mail: siddall@amnh.org

Phylogeny and systematic revision of the family Pseudokeronopsidae (Protista, Ciliophora, Hypotricha), with description of a new estuarine species of Pseudokeronopsis

Abstract: Chen, X., Clamp, J. C. & Song, W. (2011). Phylogeny and systematic revision of the family Pseudokeronopsidae (Protista, Ciliophora, Hypotricha), with description of a new estuarine species of Pseudokeronopsis. —Zoologica Scripta, 40, 659–671. The family Pseudokeronopsidae is a taxon of hypotrich ciliates with a history of uncertainty regarding its systematic and phylogenetic relationships to other members of the order Urostylida. Phylogenetic analyses of pseudokeronopsids were made using all available molecular and morphological information, and the patterns of morphogenesis of the group were reinvestigated. Results clearly demonstrated that the genera Thigmokeronopsis and Apokeronopsis are not confamilial with Pseudokeronopsis and Nothoholosticha but, instead, must be transferred to the family Urostylidae. A new estuarine species, Pseudokeronopsis erythrina sp. n. was discovered and described using a combination of morphological and molecular characters. Species of Pseudokeronopsis are difficult to characterize but can be distinguished from one another by a combination of morphological and morphogenetic characters. Pseudokeronopsis similis is a freshwater species that has significant morphogenetic differences with other members of the genus and may not be congeneric with them.

Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean

Abstract: A phylogeny is presented for the speciose, near pan-tropical, carnivorous achatinoid land-snail superfamily Streptaxoidea inferred from DNA sequences (two nuclear and two mitochondrial regions) from 114 taxa from Africa, the Indian Ocean, Asia, South America and Europe. In all analyses, Streptaxidae are monophyletic, while the (two to six) previously recognised subfamilies are polyphyletic, as are several genus-level taxa including the most speciose genus Gulella, necessitating major taxonomic review. The Asian Diapheridae are sister to Streptaxidae, which forms several well-supported clades originating in a persistent basal polytomy. Divergence dating estimates, historical biogeography, and the fossil context suggest a Cretaceous origin of these families, but suggest Gondwanan vicariance predated most radiation. The basal polytomy dates to the Paleogene and may correspond to a rapid radiation in Africa. There is evidence for multiple Cenozoic dispersals followed by radiation, including at least two from Africa to South America, at least two from Africa to Asia and at least two from Africa to Madagascar, indicating Cenozoic turnover in tropical snail faunas. The endemic Seychelles and Mascarene streptaxid faunas each are composites of early Cenozoic lineages and more recent dispersals from Africa, with no direct evidence for an Asian origin as currently proposed. Peak streptaxid diversity in East Africa is explained by Neogene speciation among a large number of coexisting ancient lineages, a phenomenon most pronounced in the Eastern Arc-Coastal Forests centre of endemism. This includes Miocene diversification in Gulella, a primarily East and South-East African group which remains strikingly diverse even after unrelated ‘Gulella’ species are reclassified.

Cryptic diversity and species assignment of large lantern sharks of the Etmopterus spinax clade from the Southern Hemisphere (Squaliformes, Etmopteridae)

Abstract: Straube, N., Kriwet, J. & Schliewen, U. K. (2010). Cryptic diversity and species assignment of large lantern sharks of the Etmopterus spinax clade from the Southern Hemisphere (Squaliformes, Etmopteridae). —Zoologica Scripta, 40, 61–75. Many species of the speciose deep-sea shark family Etmopteridae (lantern sharks) are a regular by-catch component of deepwater trawl and longline commercial fisheries. As for many elasmobranchs, the low fecundity, late sexual maturation and extreme longevity of the lantern sharks increase their susceptibility to overfishing. However, the taxonomic uncertainty within etmopterids and the poorly known patterns of dispersal of these shark species hampers the establishment of reasonable monitoring efforts. Here, we present the first molecular approach to clarify the taxonomy and distribution of a morphologically uniform group of lantern sharks comprising Etmopterus granulosus and closely related congeners by using nucleotide sequence data from the mitochondrial DNA cytochrome oxidase I gene and amplified fragment length polymorphisms. Samples were collected from several locations in the Southern Hemisphere, where the species occur. Our analyses reveal a high level of cryptic diversity. E. granulosus is not endemic to Chile, but instead has a widespread distribution in the Southern Hemisphere being synonymous to New Zealand Etmopterus baxteri. Conversely, specimens previously assigned to E. baxteri from off South Africa apparently represent a distinct species. Our results provide the basis for the re-description of E. granulosus and E. baxteri which will help in the establishment of useful monitoring and management strategies.

Molecular evolution of Cinetochilum and Sathrophilus (Protozoa, Ciliophora, Oligohymenophorea), two genera of ciliates with morphological affinities to scuticociliates

Abstract: Zhang, Q., Miao, M., Struder-Kypke, M. C., Al-Rasheid, K. A. S., Al-Farraj, S. A. & Song, W. (2011). Molecular evolution of Cinetochilum and Sathrophilus (Protozoa, Ciliophora, Oligohymenophorea), two genera of ciliates with morphological affinities to scuticociliates. —Zoologica Scripta, 40, 317–325. The ciliate order Loxocephalida sensu Li et al. (2006) has been considered to be systematically uncertain within the subclass Scuticociliatia. Loxocephalids display mixed morphological features and morphogenetic patterns that are found in two different oligohymenophorean subclasses: scuticociliates and hymenostomes. To reveal their phylogenetic positions, molecular information on this group is urgently needed but still inadequate. In the present study, we have sequenced the small subunit rRNA gene of two newly described loxocephalids, Cinetochilum ovale Gong & Song 2008; and Sathrophilus planus Fan et al. 2010; which have never been discussed based on molecular analysis. Results show: (i) all phylogenetic trees are nearly identical in placing Cinetochilum closest to the subclass Apostomatia and form a monophyletic group divergent from the typical scuticociliates, (ii) the genus Sathrophilus, together with Anoplophrya, a poorly known Astomatia, forms a peripheral branch separated from the scuticociliatian assemblage and (iii) the affiliation of the loxocephalid genera sensu Li et al. (2006) is not confirmed due to a dispersion in four deeply diverged clades. In addition, the polyphyly of the genus Cyclidium, shown in previous studies, is confirmed by our phylogenetic analyses and supported by the approximately unbiased test based on the new database in this work.

Regressive progression, progressive regression or neither? Phylogeny and evolution of the Percopsiformes (Teleostei, Paracanthopterygii)

Abstract: Dillman, C.B., Bergstrom, D.E., Noltie, D.B., Holtsford, T.P. & Mayden, R.L. (2010). Regressive progression, progressive regression or neither? Phylogeny and evolution of the Percopsiformes (Teleostei, Paracanthopterygii). —Zoologica Scripta, 40, 45–60. Cave animals have fascinated scientists for centuries, and clades consisting primarily of cave-adapted species are even more intriguing. The percopsiforms are an enigmatic group of fishes comprised of nine species in seven genera, with four species in three genera exhibiting characteristic troglomorphic features, such as a lack of pigmentation and eyes. Nucleotide characters presented here provide the first test of monophyly for both the Percopsiformes and Amblyopsidae with this character type and taxonomic completeness. Characters of ND2 support a monophyletic Percopsiformes and Amblyopsidae and further document phylogeographic subdivision in two stygobitic genera, Amblyopsis and Typhlichthys, in Amblyopsidae. Age estimates from time-calibrated branch lengths utilizing two independent intra-lineage fossils indicate that the ancestor to amblyopsids is Eocene in age, and that phylogeographic subdivision in both Amblyopsis and Typhlichthys occurred primarily in the Miocene. Interestingly, ancestral character state reconstruction for the amblyopsids strongly supports the re-evolution of eyes and body pigment. While certainly unconventional, but supported with this character set, the hypothesis provides continued challenge to Dollo’s Law.

Molecular phylogeny and systematics of Neotropical toucanets in the genus Aulacorhynchus (Aves, Ramphastidae)

Abstract: IGU¨ ENZASubmitted: 13 August 2010Accepted: 7 February 2011doi:10.1111/j.1463-6409.2011.00475.xBonaccorso, E., Guayasamin, J. M., Peterson, A. T. & Navarro-Sigu¨enza, A. G. (2011).Molecular phylogeny and systematics of Neotropical toucanets in the genus Aulacorhynchus(Aves, Ramphastidae). — Zoologica Scripta, 40, 336–349.We studied the phylogenetic relationships in the genus Aulacorhynchus, an assemblage ofNeotropical toucanets distributed from Mexico south to Bolivia. Based on mitochondrialand nuclear DNA characters, we obtained a robust hypothesis of relationships for all rec-ognized species, including good representation of distinct geographic populations. Ourresults support the monophyly of the genus Aulacorhynchus, but contradict previous taxo-nomic arrangements. The genus is made up of three major clades: the Aulacorhynchus prasi-nus complex, Aulacorhynchus huallagae + Aulacorhynchus coeruleicinctis, and Aulacorhynchushaematopygus + Aulacorhynchus sulcatus + Aulacorhynchus derbianus. Andean populations ofA. derbianus are more closely related to A. sulcatus than to Pantepuian populations ofA. derbianus, rendering A. derbianus paraphyletic. Based on the molecular phylogeny, andinformation on geographic distributions and morphological and behavioural characters, wereview the specific status of these taxa and propose a new taxonomic arrangement withinAulacorhynchus.Corresponding author: Elisa Bonaccorso, Universidad Tecnolo´gica Indoame´rica, Machala y Saba-nilla, Cotocollao, Quito, Ecuador. E-mail: elisabonaccorso@gmail.comElisa Bonaccorso, Universidad Tecnolo´gica Indoame´rica, Machala y Sabanilla, Cotocollao, QuitoEC170103, Ecuador; Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard,Lawrence, KS 66045-7561, USA. E-mail: elisabonaccorso@gmail.comJuan M. Guayasamin, Universidad Tecnolo´gica Indoame´rica, Machala y Sabanilla, Cotocollao,Quito EC170103, Ecuador. E-mail: jmguayasamin@gmail.comA. Townsend Peterson, Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard,Lawrence, KS 66045-7561, USA. E-mail: town@ku.eduAdolfo G. Navarro-Sigu¨enza, Museo de Zoologi´a ‘‘Alfonso L. Herrera,’’ Facultad de Ciencias,Universidad Nacional Auto´noma de Me´xico, Apartado Postal 70-399, Me´xico D.F. 04510, Me´xico.E-mail: adolfon@ciencias.unam.mx

Monophyly, candidate species and vicariance in Biomphalaria snails (Mollusca: Planorbidae) from the Southern Andean Altiplano

Abstract: Collado, G. A., Vila, I. & Mendez, M. A. (2011). Monophyly, candidate species and vicariance in Biomphalaria snails (Mollusca: Planorbidae) from the Southern Andean Altiplano. —Zoologica Scripta, 40, 613–622. The landscape of the Neotropical southern Andean Altiplano is characterized by a succession of closed basins originated from the Miocene to the Holocene. In this region, the number of species and phylogenetic relationships among freshwater snails of the genus Biomphalaria are uncertain. Here we obtained sequences of the mitochondrial gene cytochrome oxidase subunit I from 17 Altiplano populations, including topotypes of three nominal species, which were analyzed together with published sequences of the genus using different methods of phylogenetic reconstruction and a species of Helisoma as outgroup. The Altiplano populations conform a monophyletic group whose sister group is the Neotropical species Biomphalaria peregrina. Within this clade we recovered four main lineages well supported and congruent with geographical distributions. One clade includes topotypes restricted exclusively to the Ascotan basin, the type locality of the nominal species Biomphalaria crequii. A second clade includes sequences that correspond to topotypes restricted to the Isluga and Carcote basins, the type localities of the nominal species Biomphalaria aymara and Biomphalaria costata, respectively. Two monophyletic groups clustered snails restricted to several aquatic systems within the Caquena and Lauca basins, which may represent candidate species. The branching pattern of the sequences suggests that in the diversification of these snails, events of vicariance inferred in the Pleistocene have predominated over dispersal phenomena.

Breaking family ties: taxon sampling and molecular phylogeny of chromodorid nudibranchs (Mollusca, Gastropoda)

Abstract: Johnson, R. F. (2010). Breaking family ties: taxon sampling and molecular phylogeny of chromodorid nudibranchs (Mollusca, Gastropoda). —Zoologica Scripta, 40, 137–157. Although researchers have debated the monophyly of the diverse chromodorid nudibranchs (Chromodorididae) for over 100 years, the monophyly of this family has not been properly tested. Recent morphological and molecular phylogenetic studies have added to the debate, but have not used appropriate methods to resolve this issue. I investigate how outgroup choice and taxon sampling influences tree topology and in turn the recovery of chromodorid monophyly. As a demonstration of these potential methodological problems, I then present phylogenies resulting from different taxon-sampling schemes using the same molecular data. Taxon sampling has a strong influence on the resulting phylogenies. With comprehensive taxon sampling and outgroup selection, Cadlina is not a member of the Chromodorididae. The chromodorid nudibranchs without Cadlina are monophyletic and possibly sister to the Actinocyclidae. Additionally, I found, for the first time, support for most current family groupings in the Doridoidea. I propose a new classification in which Cadlina is not considered a member of the Chromodorididae. Instead, I resurrect the family name Cadlinidae to include the genera Cadlina and Aldisa.

Molecular and morphological phylogeny of Diplazontinae (Hymenoptera, Ichneumonidae)

Abstract: Klopfstein, S., Quicke, D. L. J., Kropf, C. & Frick, H. (2011) Molecular and morphological phylogeny of Diplazontinae (Hymenoptera, Ichneumonidae). —Zoologica Scripta, 40, 379–402. Parasitoid wasps are among the most species rich and at the same time most understudied of all metazoan taxa. To understand their diversification and test hypotheses about their evolution, we need robust phylogenetic hypotheses. Here, we reconstruct the phylogeny of the subfamily Diplazontinae using four genes and 66 morphological characters both in separate analyses and in a total evidence approach. The resulting phylogeny is highly resolved, with most clades supported by multiple independent data partitions. It contains three highly supported genus groups, for which we suggest morphological and behavioural synapomorphies. The placement of some of the genera, especially Xestopelta Dasch, is unexpected, but also supported by morphology. Most of the genera are retrieved as monophyletic, with the exception of the morphologically diverse genus Syrphoctonus Forster. We split this genus into three genera, including Fossatyloides gen. n., to restore the phylogeny–classification link. Conflict between the morphological and the molecular topology was mostly resolved in favour of the molecular partition in the total evidence approach. We discuss reasons for this finding, and suggest strategies for future taxon and character sampling in Diplazontinae.

Evolution of acoustic communication in the Gomphocerinae (Orthoptera: Caelifera: Acrididae)

Abstract: Nattier, R., Robillard, T., Amedegnato, C., Couloux, A., Cruaud, C. & Desutter-Grandcolas, L. (2011) Evolution of acoustic communication in the Gomphocerinae (Orthoptera: Caelifera: Acrididae). —Zoologica Scripta, 40, 479–497. We used a phylogenetic analysis to study the communication behaviour in the Gomphocerinae (Insecta, Orthoptera, Acrididae), which are characterised by their stridulatory apparatus and a diversified acoustic repertoire. Song-emitting behaviour, together with the general description, temporal dimension and amplitude of signals were used to describe the calling song of 34 species, resulting in a matrix of 39 acoustic characters. A phylogeny using 59 gomphocerine species and two outgroups was built based on four mitochondrial markers and the acoustic characters. Based on acoustic character optimisations, ancestral calling song was reconstructed and the evolution of calling song is discussed, with a comparison to courtship behaviours. Our results show a dynamic process of calling song evolution and underline a balance between calling and courtship diversity leading to two mating strategies corresponding to two major clades in the phylogeny.

Ontogeny and phylogeny of Metaurostylopsis cheni sp. n. (Protozoa, Ciliophora), with estimating the systematic position of Metaurostylopsis

Abstract: ONGSubmitted: 10 April 2010Accepted: 2 September 2010doi:10.1111/j.1463-6409.2010.00451.xChen, X., Huang, J. & Song, W. (2011). Ontogeny and phylogeny of Metaurostylopsis chenisp. n. (Protozoa, Ciliophora), with estimating the systematic position of Metaurostylopsis. ––Zoologica Scripta, 40, 99–111.The ciliate genus Metaurostylopsis seems to be a highly divergent marine-habiting group, ofwhich neither systematic position nor the variation of their ontogeny has been criticallychecked. In the present work, the morphology and morphogenesis during asexual divisionof a new form, Metaurostylopsis cheni sp. n., isolated from the Yellow Sea, China, were inves-tigated and comparison among known congeners was performed. The new species has twotypes of cortical granules, the larger ones of which are flattened and oval or circular in out-line with a longitudinal groove, yellow–green in colour, and arranged along the cirral rowsand dorsal kineties, whereas the smaller ones are colourless or grayish and sparsely distrib-uted. The main morphogenetic features are: (i) the entire parental ciliature, including theold oral apparatus, is renewed, (ii) the oral primordium of the proter originates de novo andbeneath the surface of the buccal cavity, that is, sub-apokinetally, (iii) the anlagen of themarginal rows and of the dorsal kineties are formed intrakinetally and (iv) fusion of themacronuclear nodules results in an irregular mass with only few branches. The small sub-unit ribosomal RNA (SSU rRNA) gene of M. cheni was sequenced. Phylogenetic analysisbased on SSU rRNA gene sequence data shows that M. cheni clusters with all other Meta-urostylopsis spp. sequenced to date indicating that the genus is monophyletic and is probablyclosely related to the Apokeronopsis–Thigmokeronopsis-group, within the order Urostylida.Corresponding author: Weibo Song, Laboratory of Protozoology, Institute of Evolution andMarine Biodiversity, Ocean University of China, Qingdao 266003, China. E-mail: wsong@ouc.edu.cnXumiao Chen and Jie Huang, Laboratory of Protozoology, Institute of Evolution and MarineBiodiversity, Ocean University of China, Qingdao 266003, China. E-mail: chenxumiao@163.com,jiehuang1228@163.com

Genetic divergence and geographic variation in the deep-water Conus orbignyi complex (Mollusca: Conoidea).

Abstract: Puillandre, N. et al. (2010) Genetic divergence and geographic variation in a deep-water cone lineage: molecular and morphological analyses of the Conus orbignyi complex (Mollusca: Conoidea).The cone snails (family Conidae) are a hyperdiverse lineage of venomous gastropods. Two standard markers, COI and ITS2, were used to define six genetically-divergent groups within a subclade of Conidae that includes Conus orbignyi; each of these was then evaluated based on their shell morphology. We conclude that three forms, previously regarded as subspecies of Conus orbignyi are distinct species, now recognized as Conus orbignyi, Conus elokismenos and Conus coriolisi. In addition, three additional species (Conus pseudorbignyi, Conus joliveti and Conus comatosa) belong to this clade. Some of the proposed species (e.g., Conus elokismenos) are possibly in turn complexes comprising multiple species. Groups such as Conidae illustrate the challenges generally faced in species delimitation in biodiverse lineages. In the case of the Conus orbignyi complex, not only are there definable, genetically divergent lineages, but also considerable geographic variation within each group. Our study suggests that an intensive analysis of multiple specimens within a single locality helps to minimize the confounding effects of geographic variation and can be a useful starting point for circumscribing different species within such a confusing complex.